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Understanding the effect of multiple infections is essential for the prediction (and eventual control) of virulence evolution. Some theoretical studies have considered the possibility that several strains coexist in the same host (coinfection), but few have taken their within-host dynamics explicitly into account. Here, we develop a nested approach based on a simple model for the interaction of parasite strains with their host's immune system. We study virulence evolution by linking the within-host dynamics to an epidemiological framework that incorporates multiple infections. Our model suggests that antigenically similar parasite strains cannot coexist in the long term inside a host. We also find that the optimal level of virulence increases with the efficiency of multiple infections. Finally, we notice that coinfections create heterogeneity in the host population (with susceptible hosts and infected hosts), which can lead to evolutionary branching in the parasite population and the emergence of a hypervirulent parasite strategy. We interpret this result as a parasite specialization to the infectious state of the hosts. Our study has experimental and theoretical implications in a virulence management perspective.

We investigate the evolution of parental care and cannibalism in a spatially structured population where adults can either help or kill juveniles in their neighborhood. We show that spatial structure can reverse the selective pressures on adult behavior, leading to the evolution of parental care, whereas the nonspatial model predicts that cannibalism is the sole evolutionary outcome. Our analysis emphasizes that evolution of such spatially structured populations is best understood at the level of the cluster of invading mutants, and we define invasion fitness as the growth rate of that cluster. We derive an analytical expression for the selective pressures on the trait and show that relatedness and Hamilton's rule are recovered as emergent properties of the spatial ecological dynamics. When adults can also help other adults, the benefits to each class of recipients are weighted by the class reproductive value, a result consistent with that of other models of kin selection. Finally, we advocate a different approach to moment equations and argue that even though the development of moment closure approximations is a necessary line of research, much-needed ecological and evolutionary insight can be gained by studying the unclosed moment equations.

Arid ecosystems are liable to undergo sudden discontinuous transitions from a vegetated to a desert state as a result of human pressure and climate change. A predictive framework about the conditions under which such transitions occur is lacking. Here, we derive and analyze a general model describing the spatial dynamics of vegetation in arid ecosystems considering local facilitation as an essential process. We investigate the conditions under which continuous or discontinuous transitions from a vegetated to a desert state are likely to occur. We focus on arid ecosystems but our approach is sufficiently general to be applied to other ecosystems with severe environmental conditions. The model exhibits bistability and vegetation patchiness. High local facilitation decreases the risk of discontinuous transitions. Moreover, for arid ecosystems where local facilitation is a driving process, vegetation patchiness indicates proximity to a transition point, but does not allow distinguishing between continuous and discontinuous transitions.

The evolution of altruism, a behaviour that benefits others at one's own fitness expense, poses a darwinian paradox. The paradox is resolved if many interactions are with related individuals so that the benefits of altruism are reaped by copies of the altruistic gene in other individuals, a mechanism called kin selection. However, recognition of altruists could provide an alternative route towards the evolution of altruism. Arguably the simplest recognition system is a conspicuous, heritable tag, such as a green beard. Despite the fact that such genes have been reported, the 'green beard effect' has often been dismissed because it is unlikely that a single gene can code for altruism and a recognizable tag. Here we model the green beard effect and find that if recognition and altruism are always inherited together, the dynamics are highly unstable, leading to the loss of altruism. In contrast, if the effect is caused by loosely coupled separate genes, altruism is facilitated through beard chromodynamics in which many beard colours co-occur. This allows altruism to persist even in weakly structured populations and implies that the green beard effect, in the form of a fluid association of altruistic traits with a recognition tag, can be much more prevalent than hitherto assumed.

Most models of virulence evolution assume that a parasite cannot raise its transmission rate without causing more harm to its host. However, the existence of such trade-off relationships has recently been challenged. Here, we study how a trade-off can emerge from a model that explicitly incorporates within-host dynamics. We find that the existence and the convexity of the trade-off are robust, which implies a definite level of evolutionarily stable virulence (ESV) for the parasite. However, we also show that the dependence of the ESV on parameter values may be very strong. One possible consequence of this sensitivity is that relationships between transmission and virulence observed across populations need not conform to the patterns expected on the basis of a common (fixed) trade-off. We discuss possible experiments and implications of our results for the development of virulence management strategies.

Temporal and spatial variations of the environment are important factors favoring the evolution of dispersal. With few exceptions, these variations have been considered to be exclusively fluctuations of habitat quality. However, since the presence of conspecifics forms part of an individual's environment, demographic stochasticity may be a component of this variability as well, in particular when local populations are small. To study this effect, we analyzed the evolution of juvenile dispersal in a metapopulation model in which habitat quality is constant in space and time but occupancy fluctuates because of demographic stochasticity. Our analysis extends previous studies in that it includes competition of resources and competition for space. Also, juvenile dispersal is not given by a fixed probability but is made conditional on the presence of free territories in a patch, whereas individuals born in full patches will always disperse. Using a combination of analytical and numerical approaches, we show that demographic stochasticity in itself may provide enough variability to favor dispersal even from patches that are not fully occupied. However, there is no simple relationship between the evolution of dispersal and various indicators of demographic stochasticity. Selected dispersal depends on all aspects of the life-history profile, including kin selection.

We investigate how the evolution of communication strategies affects signal credibility when there is common interest as well as a conflict between communicating individuals. Taking alarm calls as an example, we show that if the temptation to cheat is low, a single signal is used in the population. If the temptation increases cheaters will erode the credibility of a signal, and an honest mutant using a different signal ('a private code') will be very successful until this, in turn, is cracked by cheaters. In such a system, signal use fluctuates in time and space and hence the meaning of a given signal is not constant. When the temptation to cheat is too large, no honest communication can maintain itself in a Tower of Babel of many signals. We discuss our analysis in the light of the Green Beard mechanism for the evolution of altruism.

Most models for coevolution of hosts and parasites are based on the assumption that resistance of hosts to parasites is an all-or-nothing effect. In many cases, for example where parasites require an appropriate receptor on host cells, this is a reasonable assumption. However, in many other cases, for example where hosts mount an immune response, this picture may be too simple. An immune system is expensive to maintain, which poses a question as to how much of its resources a host should allocate to resist parasites: if the risk of infection is low, natural selection may favour hosts with less effective immune systems. As optimal allocation to defence will depend on the force of infection, and the force of infection, in turn, depends on the level of defence in the rest of the host population, a game-theoretic approach is necessary. Here I analyse a simple model for the evolution of the ability to recover from infection. If parasites are not allowed to coevolve, the outcome is a single evolutionarily stable strategy (ESS). If the parasites coevolve, multiple evolutionary outcomes are possible, one in which the parasites are relatively avirulent and common and the hosts invest little in recovery ability, and another (the escalated arms race) where parasites are rare but virulent and the hosts invest heavily in defence.